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Abstract Species' persistence in increasingly variable climates will depend on resilience against the fitness costs of environmental stochasticity. Most organisms host microbiota that shield against stressors. Here, we test the hypothesis that, by limiting exposure to temporally variable stressors, microbial symbionts reduce hosts' demographic variance. We parameterized stochastic population models using data from a 14‐year symbiont‐removal experiment including seven grass species that hostEpichloëfungal endophytes. Results provide novel evidence that symbiotic benefits arise not only through improved mean fitness, but also through dampened inter‐annual variance. Hosts with “fast” life‐history traits benefited most from symbiont‐mediated demographic buffering. Under current climate conditions, contributions of demographic buffering were modest compared to benefits to mean fitness. However, simulations of increased stochasticity amplified benefits of demographic buffering and made it the more important pathway of host–symbiont mutualism. Microbial‐mediated variance buffering is likely an important, yet cryptic, mechanism of resilience in an increasingly variable world. 

Abstract Root production influences carbon and nutrient cycles and subsidizes soil biodiversity. However, the long‐term dynamics and drivers of belowground production are poorly understood for most ecosystems. In drylands, fire, eutrophication, and precipitation regimes could affect not only root production but also how roots track interannual variability in climate.We manipulated the intra‐annual precipitation regime, soil nitrogen, and fire in four common Chihuahuan Desert ecosystem types (three grasslands and one shrubland) in New Mexico, USA, where the 100‐year record indicates both long‐term drying and increasing interannual variability in aridity. First, we evaluated how root production tracked aridity over 10–17 years using climate sensitivity functions, which quantify long‐term, nonlinear relationships between biological processes and climate. Next, we determined the degree to which perturbations by fire, nitrogen addition or intra‐annual rainfall altered the sensitivity of root production to both mean and interannual variability in aridity.All ecosystems had nonlinear climate sensitivities that predicted declines in production with increases in the interannual variance of aridity. However, root production was the most sensitive to aridity in Chihuahuan Desert shrubland, with reduced production under drier and more variable aridity.Among the perturbations, only fire altered the sensitivity of root production to aridity. Root production was more than twice as sensitive to declines with aridity following prescribed fire than in unburned conditions. Neither the intra‐annual seasonal rainfall regime nor chronic nitrogen fertilization altered the sensitivity of roots to aridity.Synthesis. Our results yield new insight into how dryland plant roots respond to climate change. Our comparison of dryland ecosystems of the northern Chihuahuan Desert predicted that root production in shrublands would be more sensitive to future climates that are drier and more variable than root production in dry grasslands. Field manipulations revealed that fire could amplify the climate sensitivity of dry grassland root production, but in contrast, the climate sensitivity of root production was largely resistant to changes in the seasonal rainfall regime or increased soil fertilization. 

ABSTRACT Disruptions to functionally important symbionts with global change will negatively impact plant fitness, with broader consequences for species' abundances, distribution, and community composition. Fungal endophytes that live inside plant leaves and roots could potentially mitigate plant heat stress from global warming. Conversely, disruptions of these symbioses could exacerbate the negative impacts of warming. To better understand the consistency and strength of warming‐induced changes to fungal endophytes, we examined fungal leaf and root endophytes in three grassland warming experiments in the US ranging from 2 to 25 years and spanning 2000 km, 12°C of mean annual temperature, and 600 mm of precipitation. We found that experimental warming disrupted symbiosis between plants and fungal endophytes. Colonization of plant tissues by septate fungi decreased in response to warming by 90% in plant leaves and 35% in roots. Warming also reduced fungal diversity and changed community composition in plant leaves, but not roots. The strength, but not direction, of warming effects on fungal endophytes varied by up to 75% among warming experiments. Finally, warming decoupled fungal endophytes from host metabolism by decreasing the correlation between endophyte community and host metabolome dissimilarity. These effects were strongest in the shorter‐term experiment, suggesting endophyte‐host metabolome function may acclimate to warming over decades. Overall, warming‐driven disruption of fungal endophyte community structure and function suggests that this symbiosis may not be a reliable mechanism to promote plant resilience and ameliorate stress responses under global change. 

Abstract Species interactions may couple the resource dynamics of different primary producers and may enhance productivity by reducing loss from the system. In low‐resource systems, this biotic control may be especially important for maintaining productivity. In drylands, the activities of vascular plants and biological soil crusts can be decoupled in space because biocrusts grow on the soil surface but plant roots are underground, and decoupled in time due to biocrusts activating with smaller precipitation events than plants. Soil fungi are hypothesized to functionally couple the plants and biocrusts by transporting nutrients. We studied whether disrupting fungi between biocrusts and plants reduces nitrogen transfer and retention and decreases primary production as predicted by the fungal loop hypothesis. Additionally, we compared varying precipitation regimes that can drive different timing and depth of biological activities.We used field mesocosms in which the potential for fungal connections between biocrusts and roots remained intact or were impeded by mesh. We imposed a precipitation regime of small, frequent or large, infrequent rain events. We used¹⁵N to track fungal‐mediated nitrogen (N) transfer. We quantified microbial carbon use efficiency and plant and biocrust production and N content.Fungal connections with biocrusts benefitted plant biomass and nutrient retention under favourable (large, infrequent) precipitation regimes but not under stressful (small, frequent) regimes, demonstrating context dependency in the fungal loop. Translocation of a¹⁵N tracer from biocrusts to roots was marginally lower when fungal connections were impeded than intact. Under large, infrequent rains, when fungal connections were intact, the C:N of leaves converged towards the C:N of biocrusts, suggesting higher N retention in the plant, and plant above‐ground biomass was greater relative to the fungal connections‐impeded treatment. Carbon use efficiency in both biocrust and rooting zone soil was less C‐limited when connections were intact than impeded, again only in the large, infrequent precipitation regime.Synthesis. Although we did not find evidence of a reciprocal transfer of C and N between plants and biocrusts, plant production was benefited by fungal connections with biocrusts under favourable conditions. 

Abstract Interactions between plants and soil microbes influence plant nutrient transformations, including nitrogen (N) fixation, nutrient mineralization, and resource exchanges through fungal networks. Physical disturbances to soils can disrupt soil microbes and associated processes that support plant and microbial productivity. In low resource drylands, biological soil crusts (“biocrusts”) occupy surface soils and house key autotrophic and diazotrophic bacteria, non‐vascular plants, or lichens. Interactions among biocrusts, plants, and fungal networks between them are hypothesized to drive carbon and nutrient dynamics; however, comparisons across ecosystems are needed to generalize how soil disturbances alter microbial communities and their contributions to N pools and transformations. To evaluate linkages among plants, fungi, and biocrusts, we disturbed all unvegetated surfaces with human foot trampling twice yearly from 2013–2019 in dry conditions in cyanobacteria‐dominated biocrusts in the Chihuahuan Desert grassland and shrubland ecosystems. After 5 years, disturbance decreased the abundances of cyanobacteria (especiallyMicrocoleus steenstrupiiclade) and N‐fixers (Scytonemasp., andSchizothrixsp.) by >77% and chlorophyllaby up to 55% but, conversely, increased soil fungal abundance by 50% compared with controls. Responses of root‐associated fungi differed between the two dominant plant species and ecosystem types, with a maximum of 80% more aseptate hyphae in disturbed than in control plots. Although disturbance did not affect¹⁵N tracer transfer from biocrusts to the dominant grass,Bouteloua eriopoda, disturbance increased available soil N by 65% in the shrubland, and decreased leaf N ofB. eriopodaby up to 16%, suggesting that, although rapid N transfer during peak production was not affected by disturbance, over the long‐term plant nutrient content was disrupted. Altogether, the shrubland may be more resilient to detrimental changes due to disturbance than grassland, and these results demonstrated that disturbances to soil microbial communities have the potential to cause substantial changes in N pools by reducing and reordering biocrust taxa. 

Abstract Regional long‐term monitoring can enhance the detection of biodiversity declines associated with climate change, improving future projections by reducing reliance on space‐for‐time substitution and increasing scalability. Rodents are diverse and important consumers in drylands, regions defined by the scarcity of water that cover 45% of Earth's land surface and face increasingly drier and more variable climates. We analyzed abundance data for 22 rodent species across grassland, shrubland, ecotone, and woodland ecosystems in the southwestern USA. Two time series (1995–2006 and 2004–2013) coincided with phases of the Pacific Decadal Oscillation (PDO), which influences drought in southwestern North America. Regionally, rodent species diversity declined 20%–35%, with greater losses during the later time period. Abundance also declined regionally, but only during 2004–2013, with losses of 5% of animals captured. During the first time series (wetter climate), plant productivity outranked climate variables as the best regional predictor of rodent abundance for 70% of taxa, whereas during the second period (drier climate), climate best explained variation in abundance for 60% of taxa. Temporal dynamics in diversity and abundance differed spatially among ecosystems, with the largest declines in woodlands and shrublands of central New Mexico and Colorado. Which species were winners or losers under increasing drought and amplified interannual variability in drought depended on ecosystem type and the phase of the PDO. Fewer taxa were significant winners (18%) than losers (30%) under drought, but the identities of winners and losers differed among ecosystems for 70% of taxa. Our results suggest that the sensitivities of rodent species to climate contributed to regional declines in diversity and abundance during 1995–2013. Whether these changes portend future declines in drought‐sensitive consumers in the southwestern USA will depend on the climate during the next major PDO cycle. 

Abstract Patterns of insect herbivory may follow predictable geographical gradients, with greater herbivory at low latitudes. However, biogeographic studies of insect herbivory often do not account for multiple abiotic factors (e.g., precipitation and soil nutrients) that could underlie gradients. We tested for latitudinal clines in insect herbivory as well as climatic, edaphic, and trait‐based drivers of herbivory. We quantified herbivory on five dominant grass species over 23 sites across the Great Plains, USA. We examined the importance of climate, edaphic factors, and traits as correlates of herbivory. Herbivory increased at low latitudes when all grass species were analyzed together and for two grass species individually, while two other grasses trended in this direction. Higher precipitation was related to more herbivory for two species but less herbivory for a different species, while higher specific root length was related to more herbivory for one species and less herbivory for a different species. Taken together, results highlight that climate and trait‐based correlates of herbivory can be highly contextual and species‐specific. Patterns of insect herbivory on dominant grasses support the hypothesis that herbivory increases toward lower latitudes, though weakly, and indicates that climate change may have species‐specific effects on plant–herbivore interactions. 

Abstract Extensive ecological research has investigated extreme climate events or long‐term changes in average climate variables, but changes in year‐to‐year (interannual) variability may also cause important biological responses, even if the mean climate is stable. The environmental stochasticity that is a hallmark of climate variability can trigger unexpected biological responses that include tipping points and state transitions, and large differences in weather between consecutive years can also propagate antecedent effects, in which current biological responses depend on responsiveness to past perturbations. However, most studies to date cannot predict ecological responses to rising variance because the study of interannual variance requires empirical platforms that generate long time series. Furthermore, the ecological consequences of increases in climate variance could depend on the mean climate in complex ways; therefore, effective ecological predictions will require determining responses to both nonstationary components of climate distributions: the mean and the variance. We introduce a new design to resolve the relative importance of, and interactions between, a drier mean climate and greater climate variance, which are dual components of ongoing climate change in the southwestern United States. The Mean × Variance Experiment (MVE) adds two novel elements to prior field infrastructure methods: (1) factorial manipulation of variance together with the climate mean and (2) the creation of realistic, stochastic precipitation regimes. Here, we demonstrate the efficacy of the experimental design, including sensor networks and PhenoCams to automate monitoring. We replicated MVE across ecosystem types at the northern edge of the Chihuahuan Desert biome as a central component of the Sevilleta Long‐Term Ecological Research Program. Soil sensors detected significant treatment effects on both the mean and interannual variability in soil moisture, and PhenoCam imagery captured change in vegetation cover. Our design advances field methods to newly compare the sensitivities of populations, communities, and ecosystem processes to climate mean × variance interactions. 

Abstract Understanding the complex and unpredictable ways ecosystems are changing and predicting the state of ecosystems and the services they will provide in the future requires coordinated, long‐term research. This paper is a product of a U.S. National Science Foundation funded Long Term Ecological Research (LTER) network synthesis effort that addressed anticipated changes in future populations and communities. Each LTER site described what their site would look like in 50 or 100 yr based on long‐term patterns and responses to global change drivers in each ecosystem. Common themes emerged and predictions were grouped into state change, connectivity, resilience, time lags, and cascading effects. Here, we report on the “state change” theme, which includes examples from the Georgia Coastal (coastal marsh), Konza Prairie (mesic grassland), Luquillo (tropical forest), Sevilleta (arid grassland), and Virginia Coastal (coastal grassland) sites. Ecological thresholds (the point at which small changes in an environmental driver can produce an abrupt and persistent state change in an ecosystem quality, property, or phenomenon) were most commonly predicted. For example, in coastal ecosystems, sea‐level rise and climate change could convert salt marsh to mangroves and coastal barrier dunes to shrub thicket. Reduced fire frequency has converted grassland to shrubland in mesic prairie, whereas overgrazing combined with drought drive shrub encroachment in arid grasslands. Lastly, tropical cloud forests are susceptible to climate‐induced changes in cloud base altitude leading to shifts in species distributions. Overall, these examples reveal that state change is a likely outcome of global environmental change across a diverse range of ecosystems and highlight the need for long‐term studies to sort out the causes and consequences of state change. The diversity of sites within the LTER network facilitates the emergence of overarching concepts about state changes as an important driver of ecosystem structure, function, services, and futures.